Three new generic records and descriptions of four new species of jumping spiders (Araneae, Salticidae) from Sri Lanka

The salticid genera Bristowia Reimoser, 1934, Habrocestum Simon, 1876 and Macaroeris Wunderlich 1992 are reported from Sri Lanka for the fi rst time. One new species of Bristowia, B. gandhii sp. nov. (♂♀), and three new species of Habrocestum, H. hantaneensis sp. nov. (♂♀), H. kodigalaensis sp. nov. (♂♀) and H. ohiyaensis sp. nov. (♂), are described and diagnosed. The male of Macaroeris nidicolens Walckenaer, 1802 is redescribed and illustrated, based on new material from Sri Lanka.


Introduction
Jumping spiders are small, diurnal predators. They capture prey by actively searching and stalking (Foelix 2011). They are highly diverse in morphology, behavior and predatory ecology (Foelix 2011;Jackson 1996;Jackson et al. 2001), which makes them attractive model organisms for the study of evolutionary phenomena. The jumping spider family (Salticidae) contains more than 595 genera and about 5838 described species placed in 7 subfamilies (Maddison 2015), making it the largest family with about 13% of all spider diversity (World Spider Catalog 2015). Currently, 64 species placed in 48 genera are known from Sri Lanka (World Spider Catalog 2015). However, recent fi eldwork and taxonomic work conducted by us suggests that this is only a fraction of the true diversity of the family on the island (Benjamin 2004(Benjamin , 2006(Benjamin , 2010(Benjamin , 2015. Here we report results of our endeavor to survey and describe this diversity, recording three new genera for the island as well as describing four new species.
The genus Bristowia Reimoser, 1934 currently comprises two species and Macaroeris Wunderlich, 1992 consists of 8 species with a widespread distribution. Ground-dwelling spiders of the genus Habrocestum Simon, 1876 are widely distributed and the genus currently contains 46 species.

R e s e a r c h a r t i c l e
urn:lsid:zoobank.org:pub:BB19E059-B45A-4665-A5A7-F4FBA53F2FB1 The aim of our present study is to report on these three new generic records (Bristowia, Habrocestum, Macaroeris) and describe four new species (B. gandhii sp. nov., H. hantaneensis sp. nov., H. kodigalaensis sp. nov. and H. ohiyaensis sp. nov). This paper is part of an ongoing island-wide study to collect and record Sri Lanka's invertebrate biodiversity.

Material and Methods
Sampling was carried out in all major climatic-physiographic zones of Sri Lanka from February 2010 to October 2015. Spiders were collected by leaf litter sifting, general hand collecting, sweeping and beating of bushes and trees. Spiders were preserved in either 70% or 100% ethanol and examined under an Olympus SZX7 or a Leica M205C stereo microscope and identifi ed morphologically to species level using available literature such as Prószyński (2003), Prószyński & Deeleman-Reinhold (2012), Żabka (1985) and several databases (Prószyński 2006(Prószyński , 2015Metzner 2015). Male palps were kept in methyl salicyclate for 4-5 hours prior to drawing. After clearing, palps were temporarily mounted on cavity slides in the same fl uid for examination and illustration. Female genitalia were excised by making holes around it using entomological pins and then the epigynal area was gently separated from the abdomen with a fi ne forceps (#5). Abdominal tissue was digested with Sigma Pancreatin LP 1750 enzyme complex, in a solution of sodium borate, prepared following the method described in Dingerkus & Uhler (1977). The specimen was then transferred to methyl salicylate and temporarily mounted for examination and illustration (Benjamin 2011). Drawings of male palps and internal structures of epigynes were made with the aid of a drawing tube attached to an Olympus BX51 microscope. Highly sclerotized or darker areas of palps and epigynum were shaded with a HB pencil. A Nikon D80 camera was used to take photographs of live spiders. Photographs of palps, epigynes and intact spiders were taken with a Leica MC170 HD camera mounted on a Leica M205C stereo microscope using Leica Application Suite, LAS version 4.6.2 software (Leica Microsystems Limited, Switzerland). Before photographing, palps and epigynes were embedded in Balea hygiene-hand gel (DM, Germany) to avoid shaking. For a single photo, between 30 and 40 images were taken in a Z-stack and merged to one image with the Helicon Focus stacking software (version 6, Helicon soft Ltd). Images were further edited with Adobe Photoshop CC and arranged in plates with Adobe Illustrator CS6. All measurements were taken with an Olympus SZX7 binocular stereo microscope with a graticule calibrated with a stage micrometer (1 mm / 100 divisions). After examination, the genitalia were transferred to micro-vials and kept in the same collection vial as the original specimen. Taxonomic histories follow the World Spider Catalog (2015). ArcGIS, version 9.3 software (Esri, USA) was used to create the GIS map for sampled localities of Sri Lanka (Fig. 1). All measurements are in millimeters. All specimens unless otherwise stated are deposited in the National Museum of Sri Lanka.  Reimoser, 1934Bristowia Reimoser, 1934, fi gs 1-3. Type species: B. heterospinosa Reimoser, 1934.

Diagnosis
Spiders of this genus are easily distinguishable by the elongated coxa, trochanter and patella of the fi rst legs, the peculiar spination of the metatarsus (2 pairs of ventral spines) and tibia I (4 prolateral and 3 retrolateral spines), and the patella and tibia with a ventral fringe of black hairs (Ono et al. 2009;Prószyński 1984;Reimoser 1934;Seo 1986;Szüts 2004). They have a longer prosoma with a thoracic fovea, and pluridentate chelicerae. The structure of the copulatory organs is relatively simple. The male palp has a single procurved retrolateral tibial apophysis, a thin embolus, and a pear-shaped tegulum. The epigynum has large, globular, spermathecae with broad insemination ducts, and a circular hollow copulatory opening (Dobroruka 2004;Ikeda 1995

Diagnosis
This species can easily be distinguished from B. afra Szüts, 2004 by the high carapace with steeper thoracic slope, a thicker and longer embolus, copulatory openings anterior to the spermathecae, a large distance between copulatory ducts and the presence of a posterior epigynal plate with a median depression, and from B. heterospinosa by broader copulatory ducts, a comparatively shallow median indentation of posterior epigynal plate and a thicker embolus.

Etymology
The species is named for Mohandas Karamchand Gandhi (1869Gandhi ( -1948. He was the pre-eminent leader of the Indian Independence Movement in British-ruled India, eventually paving the way for independence of Sri Lanka as well.

Paratype
ABDOMEN. Elongated oval-shaped and narrower than prosoma. Dorsum yellowish brown with several transverse black markings and black edges (Fig. 3A). Venter pale yellow, blackish brown near the yellowish black spinnerets.
LEGS. First pair of legs more strongly modifi ed in males than females, with elongated coxa, trochanter and patella. Patella and tibia with fringe of thick, long, black bristles (Fig. 3A). Leg I dark brown except for pale yellow tarsus, other legs yellowish brown. Tibia I with 4 prolateral and 3 retrolateral spines, metatarsus I with 2 pairs of fi ne, long spines. Legs III and IV spineless.

Distribution
India, Sri Lanka.

Diagnosis
Small spiders. Prosoma longer than abdomen and broader behind PLEs. Anterior portion fl at, extending over 75-87% of prosomal length, posterior much steeper and slightly truncated (Prószyński 2003). Short eye fi eld. Third eye row located on the edges of prosoma. Chelicerae with two teeth on promargin and single bicuspid tooth on retromargin. Abdomen rounded or oval in shape, narrower than prosoma, with a pair of large white spots and chevrons posteriorly (Prószyński 2003), fused medially in most species, dorsal view of cymbium with large, round spot of white scales (Fig. 2C). Third and fourth legs longer (tibia plus patella III ≥ tibia plus patella IV), femora III longer than others (Richman 1981). Palp with single tibial apophysis which is hook shaped in most species. Bulbus elongated. Shape of embolus species specifi c, variable in length and originating under the tegulum. Conductor present or absent (Wunderlich 2008). Epigynum with small pocket-like sclerotized vaginal roof and shallow notch at posterior edge near epigastric furrow. Copulatory openings large, located laterally along the midlength of epigyne. Large, transverse and membranous white "window", with translucent dark internal structures, very complicated consisting of large channels running anteriorly and then reversing back, passing into spermathecae with thick walls connected by soft channels (Prószyński 2003

Diagnosis
This species can be distinguished from its congeners by the cone-shaped and comparatively short embolus (Figs 5C, 6A), long tibial apophyses with serrated tip, elongated tegulum with a large proximal lobe ( Fig. 6A-B), epigyne with a wide vaginal roof, and characteristic shape of receptacles and accessory glands ( Fig. 6C-D). This species is close to the female of H. laurae Peckham & Peckham, 1903, but differs in the shape of the spermathecae and presence of accessory glands.

Etymology
The specifi c epithet refers to the type locality. COLOR AND BODY. Carapace blackish brown. Chelicerae dark brown with two promarginal teeth and single bicuspid retromarginal tooth. Labium yellowish brown with black margin. Sternum oval with black colored spots in middle, edges reddish brown. Vicinity of eyes blackish brown and slightly elevated. Front eyes covered with long dark brown bristles. First eye row slightly curved upward. Eye fi eld comparatively longer than in congeners, occupying about half the length of prosoma (Fig. 5A). Median ocular quadrangle much broader than long. White hairs interspersed on lateral sides of prosoma. Prosoma high, longer than wide. Narrow fovea in midline of prosoma behind PLEs. Lateral and posterior sides of prosoma almost vertical and posterior margin slightly truncated. Abdomen: Rounded, smaller and narrower than prosoma. Dorsum with yellowish brown small dots on anterior portion, comparably large pair of yellowish brown spots at midpoint, transverse pale yellow stripes extending laterally as dashed line at posterior half (Fig. 5A). Venter with longitudinal pale yellow dashed-line pattern from spinnerets to epigastric furrow. Spinnerets dirty yellow.

Holotype
LEGS. All legs yellow with black lateral bands. Tibia I with 3 spines on prolateral and 3 spines on retrolateral sides.  (Fig. 6D). Copulatory ducts short. Peculiarly shaped, heavily sclerotized spermathecae with thick walls. Fertilization ducts narrow arising from anterodorsal side of receptacles (Fig. 6D).  COLOR AND BODY. Carapace dark brown. Dark brown bristles near front eyes. Anterior median eyes covered with tuft of pale yellow hairs. Dark ocular region. Chelicerae blackish brown. Sternum oval shaped with dispersed black colored spots. White and brown hairs sparsely dispersed on lateral prosoma. Distance between posterior eyes slightly shorter than distance between anterior lateral eyes. Carapace somewhat high in the ocular region. Eye fi eld short and wide occupying about one-third length of prosoma. Median ocular quadrangle much broader than long. Round and middle depression around short fovea behind PLEs. Prosoma longer than wide. Lateral and posterior sides of prosoma much steeper. Posterior margin of prosoma truncated (Fig. 7A-B). Abdomen: Oval shaped, smaller and narrower than prosoma. Dorsum greyish beige color with black stripes and spots. Venter with longitudinal pale yellow dash line from spinnerets to epigastric furrow. Spinnerets blackish yellow. Anterior venter with black blotches.

Habrocestum kodigalaensis
LEGS. All legs with yellow and black banding pattern. Femora III much longer than femora IV. Front legs slightly darker than other pairs. PALP. Dark brown palp except for tibia and cymbium. Dorsum of cymbium with large, round spot of pale yellow scales. Palpal tibia with short hook-shaped tibial apophysis (Figs 7E, 8B). Elongated bulbus. Tegulum with a long lateral proximal lobe, protruding posterolaterally beyond base of cymbium. Thick embolus with hook-like tip and broader at midlength, arising from under tegular ledge (Figs 7D, 8A). Part of sperm duct clearly visible at shoulder of tegulum.

Diagnosis
This species can be distinguished from known congeners by its straight, tapering embolus ( Fig. 9B-D), medium sized, hook-shaped tibial apophysis (Fig. 9C, E) and the shape of the proximal tegular lobe (Fig. 9F). The male palp of this species is close to that of H.bovei Lucas, 1846, H. ibericum Dalmas, 1920and H. lepidum Dalmas, 1920, but it differs in the shape proximal lobe of tegulum and tegular ledge.

Etymology
The specifi c epithet refers to the type locality.

Female
Unknown.

Diagnosis
Morphologically, this genus is similar to Dendryphantes Koch, 1837, except for the structure of the genitalia (Prószyński 2003). Long prosoma broader at mid-length, fl at anterior and gently sloped posterior prosoma. Eye fi eld short and trapezoid shaped. Second row of eyes very small, rather closer to ALE than PLEs, diameter of AME twice that of ALE. Abdomen long and broad, tapering posteriorly with marginal, medial blotches and chevrons. Enlarged front legs, rather longer than other legs. Embolus long, gently curved to form a very loose coil with a wide base originating on anterolateral bulbus (Prószyński 2003;Wunderlich 2008). Tegulum distended prolaterally. Tip of tibial apophysis bent forward. Copulatory openings located antero-laterally in a depression, but not in a semilunar groove as in Dendryphantes, postero-median copulatory ducts converging diagonally, posteriorly and ending in complicated spermathecal chambers.

Diagnosis
This species is easily distinguishable from known congeners by the prosoma being broader at midlength, its massive front legs, and its broad and posteriorly tapered abdomen with an anterior white band with blotches laterally and medially and chevrons posteriorly. It has a long and gently curved embolus ( Fig. 10B-D), its tegulum is distended prolaterally and the tibial apophysis bent forward. (Merrett & Milner 2004).  COLOR AND BODY. Carapace dark reddish brown with black marking in middle of cephalic area, black colour around eyes and brown hairs surrounding AMEs. Clypeus with a few long white hairs. Chelicerae dark brown. Sparse white hairs on lateral sides of prosoma. Posterior margin of prosoma gently inclined with black colored striped pattern. Sternum dark yellow-brown. Short, trapezoid shaped eye fi eld (Fig. 10A). Cephalic region almost square and slightly elevated. Prosoma oval shaped, broader laterally behind eyes. Abdomen: long and broad in shape, tapering posteriorly. Dorsum with yellowish brown blotches and chevrons. Anterior margin with white colored net-like pattern extending laterally. Posterior margin with yellowish brown chevrons (Fig. 10A). Venter whitish grey with brown broad median longitudinal stripe from spinnerets to epigastric furrow. Spinnerets dark brown.

Material examined
LEGS. Legs I and II darker brown than the others. Front legs, strikingly massive with enlarged coxa, trochanter, femur, patella and tibia (Fig. 10A). Tibia I with 3 pairs of short stout ventral spines, tibia II with 2 pairs, metatarsi I and II with 2 pairs. Femora I-IV with 2-3 long curved dorsal spines and additionally, femora I-II with 2 short stout spines distomesally.
PALPS. Dark brown, densely clothed with pale hairs at distal end of cymbium. Retrolateral tibial apophysis strong, tip slightly bent forward, base straight (Fig. 10E). Tegulum bulbous, distended prolaterally. Long embolus with a wide basal part, originating anterolaterally on anterior part of bulbus and gently bent to form a curve that runs along depression in cymbium (Fig. 10B-D).

Discussion
Our main objective was to report on the high diversity of jumping spiders of Sri Lanka. While we added four new species and three new generic records for the island, our unpublished data, however, suggest that a huge diversity still remains to be discovered and described. We intend to provide data on the ecology and phylogenetic relationships of this unknown component of the islands' biodiversity in future contributions.